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The stomatopod crustaceans, or mantis shrimps, are colourful marine invertebrate predators. Their unusual compound eyes have dorsal and ventral regions resembling typical crustacean apposition designs separated by a unique region called the midband that consists of from two to six parallel rows of ommatidia. In species with six-row midbands, the dorsal four rows are themselves uniquely specialized for colour analysis. Rhabdoms of ommatidia in these rows are longitudinally divided into three distinct regions: an apical ultraviolet (UV) receptor, a shorter-wavelength middle tier receptor and a longer-wavelength proximal tier receptor. Each of the total of 12 photoreceptors has a different spectral sensitivity, potentially contributing to a colour-vision system with 12 channels. Mantis shrimps can discriminate both human-visible and UV colours, but with limited precision compared to other colour-vision systems. Here, we review the structure and function of stomatopod colour vision, examining the types of receptors present in a species, the spectral tuning of photoreceptors both within and across species, the neural analysis of colour and the genetics underlying the multiple visual pigments used for colour vision. Even today, after many decades of research into the colour vision of stomatopods, much of its operation and its use in nature remain a mystery. This article is part of the theme issue ‘Understanding colour vision: molecular, physiological, neuronal and behavioural studies in arthropods’.

Cephalopods have the potential to become useful experimental models in various fields of science, particularly in neuroscience, physiology, and behavior. Their complex nervous systems, intricate color- and texture-changing body patterns, and problem-solving abilities have attracted the attention of the biological research community, while the high growth rates and short life cycles of some species render them suitable for laboratory culture. Octopus chierchiae is a small octopus native to the central Pacific coast of North America whose predictable reproduction, short time to maturity, small adult size, and ability to lay multiple egg clutches (iteroparity) make this species ideally suited to laboratory culture. Here we describe novel methods for culture of O. chierchiae, with emphasis on enclosure designs, feeding regimes, and breeding management. Our results demonstrate the feasibility of multigenerational culture of O. chierchiae. Specifically, O. chierchiae bred in the laboratory grows from a 3.5-millimeter mantle length at hatching to an adult mantle length of approximately 20-30 millimeters in 250-300 days, with 14-15% survivorship to over 400 days of age in first and second generations. O. chierchiae sexually matures at around an estimated six months of age and, unlike most octopus species, can lay multiple clutches of eggs, approximately every 30-90 days. Eggs are large and hatchlings emerge as direct developing octopuses. Based on these results, we propose that O. chierchiae possesses both the practical and biological features needed for a model octopus that can be cultured repeatedly to address a wide range of fundamental biological questions.

Ahyong, Shane T., Porter, Megan L., Caldwell, Roy L. (2020): The Leopard Mantis Shrimp, Ankersquilla pardus, a New Genus and Species of Eurysquillid from Indo-West Pacific Coral Reefs. Records of the Australian Museum 72 (1): 1-8, DOI: 10.3853/j.2201-4349.72.2020.1758

Nature utilizes the available resources to construct lightweight, strong and tough materials under constrained environmental conditions. The impact surface of the fast-striking dactyl club from the mantis shrimp is an example of one such composite material; the shrimp has evolved the capability to localize damage and avoid catastrophic failure from high-speed collisions during its feeding activities. Here we report that the dactyl club of mantis shrimps contains an impact-resistant coating composed of densely packed (about 88 per cent by volume) ~65-nm bicontinuous nanoparticles of hydroxyapatite integrated within an organic matrix. These mesocrystalline hydroxyapatite nanoparticles are assembled from small, highly aligned nanocrystals. Under impacts of high strain rates (around 104 s−1), particles rotate and translate, whereas the nanocrystalline networks fracture at low-angle grain boundaries, form dislocations and undergo amorphization. The interpenetrating organic network provides additional toughening, as well as substantial damping, with a loss coefficient of around 0.02. An unusual combination of stiffness and damping is therefore achieved, outperforming many engineered materials. A coating made from densely packed hydroxyapatite particles in an organic matrix endows the dactyl club of mantis shrimps with high stiffness and energy damping.

Among marine crustaceans, biparental care has only been directly observed in the stomatopod genus Pullosquilla. This study investigated the evolutionary maintenance of biparental care in Pullosquilla thomassini (Manning, 1978) by examining fitness costs and benefits of biparental care using a manipulation of the number and sex of care providers. In the absence of parental care, egg clutch mass decreased. However, in the presence of at least one parent, there were no changes in clutch mass during the experiment. This suggests that while parental care increases production of offspring, uniparental care by either sex is sufficient. We suggest that biparental care may increase fitness in P. thomassini by facilitating the production of overlapping egg clutches. We found evidence that males providing uniparental care lost body mass, suggesting they pay a small cost in fitness. However, single males with eggs were rare at our field site (3.93% of burrows).

The predatory stomatopod crustaceans, or mantis shrimp, are among the most attractive and dynamic creatures living in the sea. Their special features include their powerful raptorial appendages, used to kill, stun, or disable other animals (whether predators, prey, or competitors), and their highly specialized compound eyes. Mantis shrimp vision is unlike that of any other animal and has several unique features. Their compound eyes are optically triple, each having three separate regions that produce overlapping visual fields viewing certain regions of space. They have the most diverse set of spectral classes of receptors ever described in animals, with as many as 16 types in a single compound eye. These receptors are based on a highly duplicated set of opsin molecules paired with strongly absorbing photostable filters in some photoreceptor types. The receptor set includes six ultraviolet types, all spectrally distinct, many themselves tuned by photostable filters. There are as many as eight types of polarization receptors of up to three spectral classes (including an ultraviolet class). In some species, two sets of these receptors analyze circularly polarized light, another unique capability. Stomatopod eyes move independently, each capable of visual field stabilization, image foveation and tracking, or scanning of image features. Stomatopods are known to recognize colors and polarization features and evidently use these in predation and communication. Altogether, mantis shrimps have perhaps the most unusual vision of any animal.

Biologically inspired solutions for modern-day sensory systems promise to deliver both higher capacity and faster, more efficient processing of information than current computational approaches. Many animals are able to perform remarkable sensing tasks despite only being able to process what would be considered modest data rates and bandwidths. The key biological innovations revolve around dedicated filter designs. By sacrificing some flexibility, specifically matched and hard-wired sensory systems, designed primarily for single roles, provide a blueprint for data and task-specific efficiency. In this paper, we examine several animal visual systems designed to use the polarization of light in spatial imaging. We investigate some implications for artificial optical processing based on models of polarization image processing in fiddler crabs, cuttlefish, octopus, and mantis shrimp.

Stomatopod crustaceans have complex visual systems containing up to 16 different spectral classes of photoreceptors, more than described for any other animal. A previous molecular study of this visual system focusing on the expression of opsin genes found many more transcripts than predicted on the basis of physiology, but was unable to fully document the expressed opsin genes responsible for this diversity. Furthermore, questions remain about how other components of phototransduction cascades are involved. This study continues prior investigations by examining the molecular function of stomatopods' visual systems using new whole eye 454 transcriptome datasets from two species, Hemisquilla californiensis and Pseudosquilla ciliata. These two species represent taxonomic diversity within the order Stomatopoda, as well as variations in the anatomy and physiology of the visual system. Using an evolutionary placement algorithm to annotate the transcriptome, we identified the presence of nine components of the stomatopods' G-protein-coupled receptor (GPCR) phototransduction cascade, including two visual arrestins, subunits of the heterotrimeric G-protein, phospholipase C, transient receptor potential channels, and opsin transcripts. The set of expressed transduction genes suggests that stomatopods utilize a Gq-mediated GPCR-signaling cascade. The most notable difference in expression between the phototransduction cascades of the two species was the number of opsin contigs recovered, with 18 contigs found in retinas of H. californiensis, and 49 contigs in those of P. ciliata. Based on phylogenetic placement and fragment overlap, these contigs were estimated to represent 14 and 33 expressed transcripts, respectively. These data expand the known opsin diversity in stomatopods to clades of arthropod opsins that are sensitive to short wavelengths and ultraviolet wavelengths and confirm the results of previous studies recovering more opsin transcripts than spectrally distinct types of photoreceptors. Many of the recovered transcripts were phylogenetically placed in an evolutionary clade of crustacean opsin sequences that is rapidly expanding as the visual systems from more species are investigated. We discuss these results in relation to the emerging pattern, particularly in crustacean visual systems, of the expression of multiple opsin transcripts in photoreceptors of the same spectral class, and even in single photoreceptor cells.

Autofluorescence is pervasive throughout the natural world and occurs in many animals, plants, and minerals. Fluorescent properties vary predictably over molt cycles of arthropod exoskeletons. This study examines intrinsic fluorescence in scorpions and crustaceans using novel fluorometric analyses and microscopy. Anchialine cave crustaceans, which we report here autofluoresce, are generally sparsely populated, and difficult for researchers to access. Therefore, little is known about the growth, molting, reproduction, and life history of their endemic inhabitants. Methods proposed here for measuring autofluorescence under near-ultraviolet wavelengths are valuable in informing techniques for optimal in situ observation and location of new larval forms, exuviae, spermataphores and eggs. We compare the intensity and emission breadth of fluorescent compounds extracted from California woodland scorpions (Uroctonus mordax) and California bay shrimp (Crangon franciscorum) to determine the effect of preservation duration on autofluorescent properties. Fluorescence data were obtained from anchialine cave crustaceans collected from Cenote Crustacea, Quintana Roo, Mexico and Dan’s Cave, Abaco Island, Bahamas. They exhibit peak emission wavelengths ranging from 429 nm to 515 nm and interspecific variation in emission peaks. Traits relating to emission spectra, such as breadth and intensity, may correlate with a variety of ecological parameters, such as sex, age, and reproductive state, allowing for potential field and laboratory-based applications.

Many biophotonic structures have their spectral properties of reflection ‘tuned’ using the (zeroth-order) Bragg criteria for phase constructive interference. This is associated with a periodicity, or distribution of periodicities, parallel to the direction of illumination. The polarization properties of these reflections are, however, typically constrained by the dimensional symmetry and intrinsic dielectric properties of the biological materials. Here we report a linearly polarizing reflector in a stomatopod crustacean that consists of 6–8 layers of hollow, ovoid vesicles with principal axes of ~550 nm, ~250 nm and ~150 nm. The reflection of unpolarized normally incident light is blue/green in colour with maximum reflectance wavelength of 520 nm and a degree of polarization greater than 0.6 over most of the visible spectrum. We demonstrate that the polarizing reflection can be explained by a resonant coupling with the first-order, in-plane, Bragg harmonics. These harmonics are associated with a distribution of periodicities perpendicular to the direction of illumination, and, due to the shape-anisotropy of the vesicles, are different for each linear polarization mode. This control and tuning of the polarization of the reflection using shape-anisotropic hollow scatterers is unlike any optical structure previously described and could provide a new design pathway for polarization-tunability in man-made photonic devices.

Some pelagic marine larvae possess anti-predator chemical defenses. Occasionally, toxic adults imbue their young with their own defensive cocktails. We examined paralarvae of the greater blue-ringed octopus (Hapalochlaena lunulata) for the deadly neurotoxin tetrodotoxin (TTX), and if present, whether TTX conferred protection to individual paralarvae. Paralarvae of H. lunulata possessed 150 ± 17 ng TTX each. These paralarvae appeared distasteful to a variety of fish and stomatopod predators, yet food items spiked with 200 ng TTX were readily consumed by predators. We conclude that TTX alone does not confer individual protection to paralarvae of H. lunulata, and that they possess an alternative defense. In larger doses, tetrodotoxin is a deterrent to the predatory stomatopod Haptosquilla trispinosa (mean dose = 3.97 μg/g). This corresponds to 12–13 paralarvae per predator based on the TTX levels of the clutch we examined. Thus, the basic assumption that individual paralarvae of H. lunulata are defended by TTX alone was disproved. Instead, functionality of TTX levels in paralarvae may arise through alternative selective pathways, such as deterrence to parasites, through kin selection, or against predator species not tested here.

Stomatopod crustaceans have complex visual systems capable of excellent spatial, colour and polarization vision. Accordingly, visual signals are used widely in their intraspecific communication behaviour. We tested the role of a particular visual signal thought to be important in mate choice in a stomatopod crustacean Haptosquilla trispinosa, by manipulating the ability of a brightly blue coloured, polarized light reflecting region on their first maxillipeds to reflect light. Depriving male H. trispinosa of their ability to produce bright blue polarized light signals did not affect their overall chances of being accepted by females. However, in comparison with control males, they had to signal significantly longer before being accepted by females, had briefer mating durations and received more aggressive responses from females before mating. These data show that males without coloured and polarized maxilliped reflections find it more difficult to be accepted by females than normal males, indicating that t...

Many organisms provision offspring with antipredator chemicals. Adult blue-ringed octopuses (Hapalochlaena spp.) harbor tetrodotoxin (TTX), which may be produced by symbiotic bacteria. Regardless of the ultimate source, we find that females invest TTX into offspring and offspring TTX levels are significantly correlated with female TTX levels. Because diversion of TTX to offspring begins during the earliest stages of egg formation, when females are still actively foraging and looking for mates, females may face an evolutionary tradeoff between provisioning larger stores of TTX in eggs and retaining that TTX for their own defense and offense (venom). Given that total TTX levels appear to increase during development and that female TTX levels correlate with those of offspring, investment may be an active adaptive process. Even after eggs have been laid, TTX levels continue to increase, suggesting that offspring or their symbionts begin producing TTX independently. The maternal investment of TTX in offspring of Hapalochlaena spp. represents a rare examination of chemical defenses, excepting ink, in cephalopods.

Stomatopod crustaceans have complex and diverse visual systems. Among their many unique features are a specialized ommatidial region (the midband) that enables the eye to have multiple overlapping visual fields, as well as sets of spectral filters that are intercalated at two levels between tiers of photoreceptors involved in polychromatic color vision. Although the physiology and visual function of stomatopod eyes have been studied for many years, how these unique visual features originated and diversified is still an open question. In order to investigate how stomatopods have attained the current complexity in visual function, we have combined physiological and morphological information (e.g. number of midband rows, number of filters in the retina, and the spectral properties of filters) with new phylogenetic analyses of relationships among species based on nucleotide sequence data from two nuclear (18S and 28S rDNA) and two mitochondrial [16S and cytochrome oxidase I (COI)] genes. Based on our recovered phylogenetic relationships among species, we propose two new superfamilies within the Stomatopoda: Hemisquilloidea and Pseudosquillodea. Maximum likelihood ancestral state reconstructions indicate that ancestral stomatopod eyes contained six midband rows and four intrarhabdomal filters, illustrating that the visual physiological complexity originated early in stomatopod evolutionary history. While the two distal filters contain conservative sets of filter pigments, the proximal filters show more spectral diversity in filter types, particularly in midband row 2, and are involved in tuning the color vision system to the photic environment. In particular, a set of related gonodactyloid families (Gonodactylidae, Protosquillidae, Takuidae) inhabiting shallow, brightly lit coral reef waters contain the largest diversity of filter pigments, which are spectrally placed relative to the underlying photoreceptors to take advantage of the broad spectrum of light available in the environment. * aLRT : approximate likelihood ratio test COI : cytochrome oxidase I GTR : general time reversible model ML : maximum likelihood PCR : polymerase chain reaction rDNA : ribosomal DNA R1–R7 : retinular cells 1–7, contained within the main rhabdom of crustacean ommatidia

The simplest prey-selection model predicts that predators select prey to maximize energetic benefit and minimize handling time. We tested whether or not energetic indices predict stomatopod preference for prey. The energetic indices included: 1) the ratio of energetic return (mass of prey tissue, M) per unit handling time (feeding duration, T); 2) the ratio of energetic benefit (caloric value of prey tissue, E) to energetic cost (caloric value of energy expended, C); and 3) the ratio of net energetic benefit (caloric value of food minus the caloric value of energy expended, NE) to handling time. Stomatopods, characterized by highly specialized raptorial feeding appendages, strike and smash the shells of prey such as snails. The prey-capture strike is one of the fastest known animal movements. Stomatopods, presented with snail shells of three size classes (5–6, 8–9, 11–12 mm in length), preferred intermediate and small snails. Oxygen consumption increased an average 2.5-fold above resting rates during feeding and increased linearly with strike frequency. The energetic cost per strike was 1.6 ± 0.2 μl O2. Stomatopods obtained more snail tissue from smashing larger snail shells. However, longer handling times were required to open shells of greater size. To obtain snail tissue, stomatopods struck larger shells a greater number of times. Strike frequency (0.72 strikes/min) was independent of shell size. The amount of snail tissue obtained per strike (0.044 mg/strike), M/T, NE/T, and E/C were all independent of snail size. The mean energetic benefit per strike (0.23 calories/strike) was 13-fold greater than the energetic cost per strike (0.017 calories/strike). Energetic cost, energetic benefit and handling time appear not to be the sole variables that explain stomatopod prey selection.

In Caribbean Panama, nonreproductive male and female stomatopods are solitary and defend their own coral-rubble cavities. When breeding pairs form, however, males assume all responsibility for cavity defense. To compare success in cavity defense and defensive tactics among paired and unpaired males, and to examine the tendency for paired stomatopods to exchange their present mates for larger (higher quality) individuals, we introduced same-sized and 15% larger male, and same-sized and 15% larger reproductive female intruders to paired and unpaired male residents in a balanced design. Paired males were more successful at cavity defense than unpaired males, evidently because paired males strike intruders more than unpaired males, and because intruders fight less intensely against paired males than against unpaired males. Paired males occasionally attempted extrapair copulations, but showed little tendency to abandon their mates in favor of larger females. Paired females, however, mated readily with intruder males that evicted resident males. Populationwide female breeding synchrony and prolonged female receptivity before oviposition reduce variance in male mating success and may force males to guard the breeding cavity to assure their paternity. Uncertainty about the reproductive condition of intruder females may prevent males from exchanging mates.

Abdopus aculeatus engages in frequent aggression and copulation, exhibits male mate-choice, and employs multiple mating tactics. Here we draw upon established hypotheses to compare male-male aggression (MMA) and male-female aggression (MFA), as they relate to their mating behavior in the wild. When contesting for females, males appear to balance mate preference (resource value) with perceived chances of winning contests (resource holding potential). Although males spent more time mating with and contesting for large "Adjacent Guarded" females (those occupying a den within arm's reach of a large "Adjacent Guarding" male), they exhibited higher rates of aggression over nonadjacent "Temporarily Guarded" females that may be more accessible. The major determinant of male-male aggressive success was size, and this factor may dictate the expression of conditional mating tactics in males. "Adjacent Guarding" males were the largest and most aggressively successful males, earning the most time copulating with females. They are considered to have the highest resource holding potential (RHP) in MMA. By contrast, in MFA, some larger individuals fled from smaller individuals, indicating that RHP appears to be a function of both size and sex in intersexual aggression. This result suggests variation in aggressiveness, or potential for severe injury-even sexual cannibalism during MFA. Male-female aggression may also be influenced by the sexual nonreceptivity of some individuals, or attempts by both sexes to increase foraging behavior by delaying mate-guarding activity.

In-depth studies on the intra-organismal distribution of toxin may yield valuable clues about potential ecological functions. The distribution of tetrodotoxin (TTX) in previously unexamined tissues of two species of blue-ringed octopuses, wild-caught Hapalochlaena fasciata and Hapalochlaena lunulata from the aquarium industry, was surveyed. Tissues from each individual were examined separately. Tetrodotoxin was detected in posterior salivary gland (PSG), arm, mantle, anterior salivary glands, digestive gland, testes contents, brachial heart, nephridia, gill, and oviducal gland of H. fasciata. By contrast TTX was found only in the PSG, mantle tissue, and ink of H. lunulata. The highest concentrations of TTX resided in the PSG of both species; however, the arms and mantle contained the greatest absolute amounts of TTX. Minimum total amounts of TTX per octopus ranged from 60 to 405 microg in H. fasciata and from 0 to 174 microg in H. lunulata and correlated well with the amounts in the PSG. Transport of TTX in the blood is loosely suggested by the presence of the toxin in blood-rich organs such as the gill and brachial hearts. The distributional data also suggest both offensive and defensive functions of TTX.

Stomatopod crustaceans (mantis shrimp) are aggressive benthic marine predators with extraordinary color vision. When communicating with conspecifics, many stomatopods display conspicuously colored body areas, often in combination with other types of signals such as motion and chemical cues. Some species occupy wide depth ranges (>30 m), where changing light conditions can influence color perception. To test the potential effects of differing ambient lights on signaling behavior, stomatopods (Gonodactylus smithii) interacted with conspecifics in aquaria, under full-spectrum, high intensity light or light restricted in either spectrum or intensity. During intrasexual and intersexual trials in full-spectrum, high intensity light, animals performed more aggressive acts using colored body parts (meral spread, lunge, strike). Stomatopods used significantly more antennular flicking, and performed aggressive acts at reduced distances under restricted light conditions. To compare the use of antennules in visual an...

Visual behaviors are prominent components of intra- and interspecific communication in shallow-water cephalopods. Meso- and bathypelagic cephalopods were believed to have limited visual communication, other than bioluminescence, due to the reduced illumination at depth. To explore potential visual behaviors in mesopelagic squid, we used undersea vehicles to observe 76 individuals of Octopoteuthis deletron. In contrast to predictions, we found this species capable of a variety of visually linked behaviors not previously reported for a deep-ocean cephalopod. The resultant ethogram describes numerous chromatic, postural, locomotor, and bioluminescent behavioral components. A few common body patterns-the whole appearance of the individual involving multiple components-are characterized. The behaviors observed from individual squid were compared using a Non-metric Multi-Dimensional Scaling (NMDS) ordination, onto which hydrographic and observation parameters were mapped. Observation length, specimen collection, and contact with the vehicle affected which behaviors were performed. A separate NMDS, analyzing the body patterns, indicated that these sets of behavioral components could be visualized as groups within the NMDS ordination. While the functional roles of the behaviors described are not yet known, our findings of numerous behaviors in O. deletron clearly indicate that bioluminescence is not the sole method of visual communication by deep-sea squid.

SummaryWe describe the addition of a fourth visual modality in the animal kingdom, the perception of circular polarized light. Animals are sensitive to various characteristics of light, such as intensity, color, and linear polarization [1, 2]. This latter capability can be used for object identification, contrast enhancement, navigation, and communication through polarizing reflections [2–4]. Circularly polarized reflections from a few animal species have also been known for some time [5, 6]. Although optically interesting [7, 8], their signal function or use (if any) was obscure because no visual system was known to detect circularly polarized light. Here, in stomatopod crustaceans, we describe for the first time a visual system capable of detecting and analyzing circularly polarized light. Four lines of evidence—behavior, electrophysiology, optical anatomy, and details of signal design—are presented to describe this new visual function. We suggest that this remarkable ability mediates sexual signaling and mate choice, although other potential functions of circular polarization vision, such as enhanced contrast in turbid environments, are also possible [7, 8]. The ability to differentiate the handedness of circularly polarized light, a visual feat never expected in the animal kingdom, is demonstrated behaviorally here for the first time.

The mating system of Abdopus aculeatus incorporates sneaker matings, mate guarding, sex-specific body patterns, frequent copulations, and male–male competition for mates, making it more similar to that of aggregating decapod cephalopods than any previously known octopus social system. Large male–female A. aculeatus occupy ‘Adjacent’ (GA) dens and copulate frequently in mate-guarding situations over successive days. Nearby individuals copulate in ‘Temporary guarding’ (GT) and ‘Transient’ (T; non-guarding) situations, the latter of which can involve ‘Sneaker’ (S) mating. In a focal animal study of these octopuses in the wild (Sulawesi, Indonesia) we addressed the hypotheses that they demonstrate: (1) precopulatory mate choice, (2) differential copulation rates by individuals employing different mating tactics, and (3) distant sex identification. We quantified daily copulation rates of A. aculeatus of reproductive size as well as aspects of copulation duration, display, mate-competition, and mate rejection. Mating tactic correlated with daily copulation rates. ♂GA spent significantly more time copulating than did ♂T, while ♀GA spent more than twice as much time per day in copula than did other females. Sneaker copulations lasted longer than those by males adopting other tactics. Mate-guarding was an effective and important tactic used by males to temporarily monopolize mating with apparently non-selective females. Males demonstrated clear pre-copulatory mate choice by guarding and mating repeatedly with large females (typically ♀GA). While foraging alone away from the den, ♂G procured ‘Transient’ copulations with unguarded females. However, mate-guarding reduced the amount of time ♂G were alone and may impede their ability to seek out new mates. Low-copulation rates by ♀T, the smallest female tactic on average, may reflect this trade-off between mate preference and mate-searching by males, or non-receptivity of some females. A male-typical body pattern (black and white stripes) appeared to facilitate distant sex identification. Although mating and aggression were often initiated before contact between individuals, same-sex copulations and intense male–female aggression were rare. By contrast frequent male–female copulations and intense male–male aggression were consistent behavioral components of mating in A. aculeatus at these sites. Because the behavioral and ecological characters conducive to this complex system are not exclusive to A. aculeatus, it is possible that other octopuses exhibit some or all of these behaviors.

SUMMARYMantis shrimp (Stomatopoda) generate extremely rapid and forceful predatory strikes through a suite of structural modifications of their raptorial appendages. Here we examine the key morphological and kinematic components of the raptorial strike that amplify the power output of the underlying muscle contractions. Morphological analyses of joint mechanics are integrated with CT scans of mineralization patterns and kinematic analyses toward the goal of understanding the mechanical basis of linkage dynamics and strike performance. We test whether a four-bar linkage mechanism amplifies rotation in this system and find that the rotational amplification is approximately two times the input rotation, thereby amplifying the velocity and acceleration of the strike. The four-bar model is generally supported, although the observed kinematic transmission is lower than predicted by the four-bar model. The results of the morphological, kinematic and mechanical analyses suggest a multi-faceted mechanical system that integrates latches, linkages and lever arms and is powered by multiple sites of cuticular energy storage. Through reorganization of joint architecture and asymmetric distribution of mineralized cuticle, the mantis shrimp's raptorial appendage offers a remarkable example of how structural and mechanical modifications can yield power amplification sufficient to produce speeds and forces at the outer known limits of biological systems.

Many stomatopod species seem capable of individual recognition. This ability appears most often in species that face severe competition for shelter, or that create shelters that are costly to reproduce. Mantis shrimp identify specific individuals (conspecific or otherwise), and adapt their defensive or offensive strategies in response to previous encounters with that opponent. Stomatopods also use individual recognition in reproductive contexts: to recognize current mates and young, and to avoid previous mates. Current thinking is that individual recognition serves to limit lethal aggression, always a risk due to the legendary strikes of their powerful raptorial appendages. The most aggressive species, with the most complex behavioral repertoires, appear to be most capable in this arena. This chapter describes the well-developed visual and chemical senses and the learning that supports this survival strategy, and then focuses on the evidence supporting chemically-mediated individual recognition. This is followed by accounts of the roles played by visual and auditory cues in individual recognition.

Stomatopods (mantis shrimp), numbering over 450 species, are renowned for their exceptional visual and chemosensory abilities and yet virtually nothing is known about their use of acoustic signals. We present acoustic recordings and analyses of the sounds of a stomatopod, Hemisquilla californiensis. This species generates tonal, low frequency sounds, which we term ‘rumble’, that are spectrally similar to those produced by African and Asian elephants. The fundamental frequency of the stomatopod rumble ranges from 20 to 60 Hz, with a strong second harmonic. Hemisquilla californiensis appears to generate these sounds through contractions of the posterior mandibular remotor muscle which is coupled to a stiff, lateral extension of the carapace. The sides of the carapace are covered by large, polarized, red spots which vibrate during sound production. Thus, the animals may generate a multi-modal signal by coupling vibrational and visual signals. Hemisquilla californiensis generates the rumble while interacting ...

SUMMARY Mantis shrimp are renowned for their unusual method of breaking shells with brief, powerful strikes of their raptorial appendages. Due to the extreme speeds of these strikes underwater, cavitation occurs between their appendages and hard-shelled prey. Here we examine the magnitude and relative contribution of the impact and cavitation forces generated by the peacock mantis shrimp Odontodactylus scyllarus. We present the surprising finding that each strike generates two brief, high-amplitude force peaks, typically 390–480 μs apart. Based on high-speed imaging, force measurements and acoustic analyses, it is evident that the first force peak is caused by the limb's impact and the second force peak is due to the collapse of cavitation bubbles. Peak limb impact forces range from 400 to 1501 N and peak cavitation forces reach 504 N. Despite their small size, O. scyllarus can generate impact forces thousands of times their body weight. Furthermore, on average, cavitation peak forces are 50% of the limb's impact force, although cavitation forces may exceed the limb impact forces by up to 280%. The rapid succession of high peak forces used by mantis shrimp suggests that mantis shrimp use a potent combination of cavitation forces and extraordinarily high impact forces to fracture shells. The stomatopod's hammer is fundamentally different from typical shell-crushing mechanisms such as fish jaws and lobster claws, and may have played an important and as yet unexamined role in the evolution of shell form.

Illegal fishing with explosives has damaged coral reefs throughout Southeast Asia. In addition to killing fish and other organisms, the blasts shatter coral skeletons, leaving fields of broken rubble that shift in the current, abrading or burying new coral recruits, and thereby slowing or preventing reef recovery. Successful restoration and rehabilitation efforts can contribute to coral reef conservation. We used field experiments to assess the effectiveness of different low-cost methods for coral reef rehabilitation in Komodo National Park (KNP), Indonesia. Our experiments were conducted at three different spatial scales. At a scale of 1 × 1m plots, we tested three different rehabilitation methods: rock piles, cement slabs, and netting pinned to the rubble. Significantly more corals per square meter grew on rocks, followed by cement, netting, and untreated rubble, although many plots were scattered by strong water current or buried by rubble after 2.5 years. To test the benefits of the most successful treatment, rocks, at more realistic scales, we established 10 × 10 m plots of rock piles at each of our nine sites in 2000. Three years after installation, coverage by hard corals on the rocks continued to increase, although rehabilitation in high current areas remained the most difficult. In 2002 rehabilitation efforts in KNP were increased over 6000 m 2 to test four rock pile designs at each of four rubble field sites. Assuming that there is an adequate larval supply, using rocks for simple, low-budget, large-scale rehabilitation appears to be a viable option for restoring the structural foundation of damaged reefs.

There have been few studies that demonstrate a protective function of inking behavior of cephalopods. In this paper I report the use of ink pseudomorphs by adult Octopus bocki against predatory attacks from green turtle (Chelonia mydas) hatchlings. Turtles that attacked ink pseudomorphs ceased predation attempts whereas naive turtles attacked and ingested octopuses.

We have generated a BAC library from the Indonesian coelacanth, Latimeria menadoensis. This library was generated using genomic DNA of nuclei isolated from heart tissue, and has an average insert size of 171 kb. There are a total of 288 384-well microtiter dishes in the library (110,592 clones) and its genomic representation is estimated to encompass ≥7X coverage based on the amount of DNA presumably cloned in the library as well as via hybridization with probes to a small set of single copy genes. This genomic resource has been made available to the public and should prove useful to the scientific community for many applications, including comparative genomics, molecular evolution and conservation genetics. J. Exp. Zool. 301A:228–234, 2004. © 2004 Wiley-Liss, Inc.

Visual pigments, the molecules in photoreceptors that initiate the process of vision, are inherently dichroic, differentially absorbing light according to its axis of polarization. Many animals have taken advantage of this property to build receptor systems capable of analyzing the polarization of incoming light, as polarized light is abundant in natural scenes (commonly being produced by scattering or reflection). Such polarization sensitivity has long been associated with behavioral tasks like orientation or navigation. However, only recently have we become aware that it can be incorporated into a high-level visual perception akin to color vision, permitting segmentation of a viewed scene into regions that differ in their polarization. By analogy to color vision, we call this capacity polarization vision. It is apparently used for tasks like those that color vision specializes in: contrast enhancement, camouflage breaking, object recognition, and signal detection and discrimination. While color is very useful in terrestrial or shallow-water environments, it is an unreliable cue deeper in water due to the spectral modification of light as it travels through water of various depths or of varying optical quality. Here, polarization vision has special utility and consequently has evolved in numerous marine species, as well as at least one terrestrial animal. In this review, we consider recent findings concerning polarization vision and its significance in biological signaling.

Visual pigments in many animal species, including stomatopod crustaceans, are adapted to the photic environments inhabited by that species. However, some species occupy a diversity of environments as adults (such as a range of depths in the ocean), and a single set of visual pigments would not be equally adaptive for all habitats in which individuals live. We characterized the visual pigment complements of three species of stomatopod crustaceans, Haptosquilla trispinosa, Gonodactylellus affinis, and Gonodactylopsis spongicola, which are unusual for this group in that each lives at depths from the subtidal to several tens of meters. Using microspectrophotometry, we determined the visual pigments in all classes of main rhabdoms in individuals of each species from shallow or deep habitats. Each species expressed the typical diversity of visual pigments commonly found in stomatopods, but there was little or no evidence of differential expression of visual pigments in animals of any species collected from different depths. Vision in these species, therefore, is not tuned to spectral characteristics of the photic environment by varying the assemblages of visual pigments appearing in their retinas.

The ‘disco’ or ‘electric’ clam Ctenoides ales (Limidae) is the only species of bivalve known to have a behaviourally mediated photic display. This display is so vivid that it has been repeatedly confused for bioluminescence, but it is actually the result of scattered light. The flashing occurs on the mantle lip, where electron microscopy revealed two distinct tissue sides: one highly scattering side that contains dense aggregations of spheres composed of silica, and one highly absorbing side that does not. High-speed video confirmed that the two sides act in concert to alternate between vivid broadband reflectance and strong absorption in the blue region of the spectrum. Optical modelling suggests that the diameter of the spheres is nearly optimal for scattering visible light, especially at shorter wavelengths which predominate in their environment. This simple mechanism produces a striking optical effect that may function as a signal.

The compound eyes of mantis shrimps (stomatopod crustaceans) include an unparalleled diversity of visual pigments and spectral receptor classes in retinas of each species. We compared the visual pigment and spectral receptor classes of 12 species of gonodactyloid stomatopods from a variety of photic environments, from intertidal to deep water (> 50 m), to learn how spectral tuning in the different photoreceptor types is modified within different photic environments. Results show that receptors of the peripheral photoreceptors, those outside the midband which are responsible for standard visual tasks such as spatial vision and motion detection, reveal the well–known pattern of decreasing λ max with increasing depth. Receptors of midband rows 5 and 6, which are specialized for polarization vision, are similar in all species, having visual λ max –values near 500 nm, independent of depth. Finally, the spectral receptors of midband rows 1 to 4 are tuned for maximum coverage of the spectrum of irradiance available in the habitat of each species. The quality of the visual worlds experienced by each species we studied must vary considerably, but all appear to exploit the full capabilities offered by their complex visual systems.

A detailed account of the second coelacanth specimen known to have been captured in Indonesian waters is provided. The account is written in the official format of the Coelacanth Conservation Council's (CCC) inventory of known specimens of the living coelacanth Latimeria chalumnae, and the specimen is herein numbered CCC no. 175. Morphological measurements and fin ray counts are recorded. A comparison of these measurements and meristic data with those recorded for Comoran specimens of Latimeria chalumnae shows that the present specimen has a similar allometry to L. chalumnae. Ongoing genetic analyses should reveal the depth of divergence between the two known populations of living coelacanths and help determine if this specimen is conspecific with its Comoran counterparts.

We examined the visual systems of two species of the lysiosquilloid stomatopod crustacean genus Pullosquilla living sympatrically in waters around the island of Moorea, French Polynesia. P. litoralis lives in bright, shallow environments to a maximum depth of less than 2 m, while P. thomassini occupies depths to at least 37m. By occupying different environments in the same geographic regions, the closely related species pair affords a rare opportunity to examine the effect of photic environment on visual evolutionary adaptation. We quantified retinal anatomy, determined the spectral diversity of visual pigments, and measured the absorption spectra of intrarhabdomal filters in retinas of both species to compare their complements of spectral receptors. The two species were similar in most respects, but the visual pigment in the main retina of P. thomassini had λmax at 467 nm, compared to 509 nm in the corresponding retinal region of P. litoralis. The absorption of visual pigments in Row 3 of the midband, th...

Larvae of decapod and stomatopod crustaceans possess paired compound eyes not unlike those of adult crustaceans. However, the visual demands of larval and adult life differ considerably. Furthermore, the eyes of adult stomatopods appear to be far more specialized than those of the larvae. We examined eyes of several stomatopod species just before and after larval metamorphosis. At this time, the entire larval retina is joined by a new, adult-type retinal array which gradually replaces the remnants of the larval retina. The new retina of the postlarva is anatomically similar to that of the full-grown adult, and has virtually identical assemblages of intrarhabdomal filters. We determined the photopigments of Gonodactylus aloha, the only species for which we were able to obtain both larval and adult specimens, using microspectrophotometry. The single middle-wavelength larval rhodopsin (λmax= 499 nm) disappears at metamorphosis; none of the 10 classes of adult rhodopsins has λmax between 473 and 510 nm. This metamorphic change of visual pigment does not occur in a comparison species of decapod crustacean, the blue crab Callinectes sapidus. Here, rhodopsins both of the megalops larva and the adult had λmax at 503-504 nm. The difference between these two species can be explained by the varying ecological requirements of their larvae and adults, and more study of visual pigments in retinas of larval and adult crustaceans is warranted.

Blue-ringed octopuses (genus Hapalochlaena) possess the potent neurotoxin tetrodotoxin (TTX). We examined the microdistribution of TTX in ten tissues of Hapalochlaena lunulata and Hapalochlaena fasciata by immunolabeling for fluorescent light microscopy (FLM). We visualized TTX throughout the posterior salivary gland, but the toxin was concentrated in cells lining the secretory tubules within the gland. Tetrodotoxin was present just beneath the epidermis of the integument (mantle and arms) and also concentrated in channels running through the dermis. This was suggestive of a TTX transport mechanism in the blood of the octopus, which would also explain the presence of the toxin in the blood-rich brachial hearts, gills, nephridia, and highly vascularized Needham's sac (testes contents). We also present the first report of TTX in any cephalopod outside of the genus Hapalochlaena. A specimen of Octopus bocki from French Polynesia contained a small amount of TTX in the digestive gland.